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2020년 2월 박사학위 논문
한국산 등각류의 계통분류학적 연구
A Systematic Study of the Isopods (Crustacea, Peracarida) from Korea
조선대학교 대학원
생 명 과 학 과
김 성 훈
[UCI]I804:24011-200000288711
[UCI]I804:24011-200000288711
한국산 등각류의 계통분류학적 연구
A Systematic Study of the Isopods (Crustacea, Peracarida) from Korea
2020년 2월 25일
조선대학교 대학원
생 명 과 학 과
김 성 훈
한국산 등각류의 계통분류학적 연구
지도교수 윤 성 명
이 논문을 이학 박사학위 신청 논문으로 제출함 2019년 10월
조선대학교 대학원
생 명 과 학 과
김 성 훈
김성훈의 박사학위논문을 인준함
위 원 장 경북대학교 교 수 황의욱 (인) 위 원 국립생태원 선 임 연 구 원 정태원 (인) 위 원 조선대학교 교 수 이정섭 (인) 위 원 조선대학교 교 수 전택중 (인) 위 원 조선대학교 교 수 윤성명 (인)
2019년 12월
조선대학교 대학교
i
CONTENTS
LIST OF FIGURES ··· iii
LIST OF TABLES ··· xiii
국문초록 ··· xiv
Part 1. Morphological Study of the Marine Isopods (Crustacea, Peracarida) from Korea ··· 1
1. Introduction ··· 2
2. Materials and methods ··· 4
3. Morphology and taxonomic characters ··· 6
3.1. General morphology ··· 6
4. Systematic accounts ··· 13
4.1. List of species ··· 13
4.2. Descriptions of species ··· 19
5. Reference ··· 236
ii
Part 2. Cladistic analysis of the Isopods ··· 248
1. Introduction ··· 249
2. Materials and methods ··· 251
2.1. Data sources ··· 251
2.2. Characters and character states ··· 254
2.3. Cladistic analysis ··· 266
3. Results ··· 267
3.1. Cladograms ··· 267
3.2. Character transition in the cladograms ··· 268
4. Discussion ··· 277
5. Reference ··· 283
ABSTRACT ··· 287
APPENDIX ··· 289
iii
LIST OF FIGURES
Fig. 1. Localities of the present study (the list is provided in Appendix 1). ··· 5 Fig. 2. General morphology of body. A, Habitus, dorsal view (left, Metacirolana; right, Janiralata);
B, Habitus, lateral view (Metacirolana). ··· 9 Fig. 3. General morphologies of cephalon and cephalic appendages. A, Antennule (Chaetiliidae); B, Antenna (Janiridae); C, Ventral view of cephalon (Cirolanidae); D, Mandible (Idoteidae); E, Mandible (Cirolanidae); F, Maxillule (Cirolanidae); G, Maxilla (Cirolanidae); H, Maxilliped (Idoteidae). ··· 10 Fig. 4. Morphologies of pereopods. A, Prehensile pereopod (Cymothoidae); B, Subchelate pereopod (Anthuridae); C, Ambulatory pereopod (Cirolanidae); D, Natatory pereopod (Cirolanidae); E Pylopod (Gnathiidae). ··· 11 Fig. 5. Morphologies of pleonal appendages. A, Pleopod 1 (Anthuridae); B, Pleopod 2 of male (Idoteidae); C, Uropod (Janiridae); D, Uropod (Cirolanidae); E, Uropod (Idoteidae); F, Uropod (Anthuridae); G, Pleotelson (Anthuridae). ··· 12 Fig. 6. Amakusanthura n. sp., holotype, female. A, Habitus, dorsal view; B, Habitus, lateral view; C, Antennule; D, Antenna; E, Mandible; F, Maxilla; G, Maxilliped. Scale bars: A, B = 1 mm, C−G
= 0.1 mm. ··· 23 Fig. 7. Amakusanthura n. sp., holotype, female. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar = 0.1 mm. ··· 25 Fig. 8. Amakusanthura n. sp., holotype, female. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F, Uropodal exopod; G, Uropodal endopod; H, Pleotelson. Scale bar:
A−H = 0.5 mm. ··· 27 Fig. 9. Apanthura n. sp., holotype, female. A, Habitus, dorsal view; B, Habitus, lateral view; C, Antennule; D, Antenna; E, Mandible; F, Maxilla; G, Maxilliped. Scale bars: A, B = 1 mm, C−G
= 0.1 mm. ··· 30 Fig. 10. Apanthura n. sp., holotype, female. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 0.1 mm. ··· 32
iv
Fig. 11. Apanthura n. sp., holotype, female. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F, Uropodal exopod; G, Uropodal endopod; H, Pleotelson. Scale bar: A−H=0.1 mm. ··· 34 Fig. 12. Caenanthura engimatica, female. A, Habitus, dorsal view; B, Habitus, lateral view; C, Pleotelson; D, Antennule; E, Antenna. Scale bars: A, B = 1 mm; C−E = 0.2 mm. (cited from Kim & Yoon, 2018a) ··· 37 Fig. 13. Caenanthura engimatica, female. A, Right mandible; B, Left mandible; C, Maxilla; D, Maxilliped. Scale bar: A−D = 0.2 mm. (cited from Kim & Yoon, 2018a) ··· 39 Fig. 14. Caenanthura engimatica, female. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G= 0.5 mm. (cited from Kim & Yoon, 2018a) ··· 41 Fig. 15. Caenanthura engimatica, female. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4;
E, Pleopod 5; F, Uropodal exopod; G, Uropodal endopod. Scale bar: A−G = 0.5 mm. (cited from Kim & Yoon, 2018a) ··· 43 Fig. 16. Kupellonura n. sp. 1, holotype, female. A, Habitus, dorsal view; B, Pleotelson, dorsal view;
C, Antennule; D, Antenna; E, Mandible; F, Maxilla; G, Maxilliped. Scale bars: A = 1 mm, B−G
= 0.2 mm. ··· 47 Fig. 17. Kupellonura n. sp. 1, holotype, female. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 0.2 mm. ··· 49 Fig. 18. Kupellonura n. sp. 1, holotype, female. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F, Uropodal exopod; G, Uropodal endopod. Scale bar: A−G = 0.2 mm.
··· 51 Fig. 19. Kupellonura n. sp. 2, holotype, female. A, Habitus, dorsal view; B, Pleotelson and uropod, dorsal view; C, Antennule; D, Antenna; E, Mandible; F, Maxilla; G, Maxilliped. Scale bars: A
= 1 mm, B−D = 0.2 mm, E−G = 0.05 mm. ··· 53 Fig. 20. Kupellonura n. sp. 2, holotype, female. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 0.2 mm. ··· 55
v
Fig. 21. Kupellonura n. sp. 2, holotype, female. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F, Uropodal exopod; G, Uropodal endopod. Scale bars: A−G = 0.2 mm. ··· 57 Fig. 22. Eurydice longiantenata. A, Habitus, dorsal view; B, Habitus, lateral view. Scale bar: A, B
= 2 mm. (cited from Kim & Yoon, 2019a) ··· 66 Fig. 23. Eurydice longiantenata, male. A, Habitus, dorsal view; B, Habitus, lateral view; C, Cephalon, ventral view; D, Pleotelson; E, Antennule; F, Antenna; G, Proximal region of antenna; H, Distal end of antenna; I, Penes. Scale bars: A, B = 1 mm, C, D = 0.5 mm, E−I = 0.2 mm. (cited from Kim & Yoon, 2019a) ··· 68 Fig. 24. Eurydice longiantenata, male. A, Right mandible; B, Left mandible; C, Maxillule, D, Maxilla; E, Maxilliped. Scale bars: A−E = 0.5 mm. (cited from Kim & Yoon, 2019a) ··· 70 Fig. 25. Eurydice longiantenata, male. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 0.5 mm. (cited from Kim &
Yoon, 2019a) ··· 72 Fig. 26. Eurydice longiantenata, male. A, Pleopod 1; B, Pleopod 2; C, Appendix masculina; D, Pleopod 3; E, Pleopod 4; F, Pleopod 5; G, Uropod; H, Distal region of uropodal endopod; I, Distal region of uropodal exopod. Scale bars: A, B, D−G = 0.5 mm, C, H, I = 0.1 mm. (cited from Kim & Yoon, 2019a) ··· 73 Fig. 27. Metacirolana japonica. A, Habitus, male; B, Habitus, female. Scale bar: A, B = 1 mm. (cited from Kim & Yoon, 2019a) ··· 77 Fig. 28. Metacirolana japonica, male (A, B, D−F) and female (C). A, Habitus, dorsal view; B, Habitus, lateral view; C, Habitus, dorsal view; D, Cephalon, anterodorsal view; E, Cephalon, ventral view; F, Pleotelson; G, Penes. Scale bars: A−C = 1 mm, D−G = 0.2 mm. (cited from Kim & Yoon, 2019a) ··· 79 Fig. 29. Metacirolana japonica, male. A, Antennule; B, Antenna; C, Right mandible; D, Left mandible; E, Maxillule, F, Maxilla; G, Maxilliped. Scale bars: A−G = 0.1 mm. (cited from Kim
& Yoon, 2019a) ··· 81
vi
Fig. 30. Metacirolana japonica, male. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 0.5 mm. (cited from Kim &
Yoon, 2019a) ··· 82 Fig. 31. Metacirolana japonica, male. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F, Uropod. Scale bar: A−F = 0.5 mm. (cited from Kim & Yoon, 2019a) ··· 84 Fig. 32. Nerocila sp. A, Habitus, dorsal view; B, Habitus, lateral view; C, Cephalon, dorsal view; D, Cephalon, ventral view; E, Pleotelson, dorsal view; F, Distal end of pleotelson. Scale bars: A, B = 1 cm, C−F = 1 mm. ··· 88 Fig. 33. Nerocila sp. A, Antennule; B, Antenna; C, Left mandible; D, Right mandible; E, Maxillule;
F, Distal end of maxillule; G, Maxilla; H, Distal end of maxilla; I, Maxilliped. Scale bars: A−D
= 0.4 mm, E, G, I = 0.2 mm, F, H = 0.05 mm. ··· 90 Fig. 34. Nerocila sp. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5;
F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 1 mm. ··· 91 Fig. 35. Nerocila sp. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F, Uropod. Scale bar: A−F = 1 mm. ··· 93 Fig. 36. Elaphognathia kikuchii, male. A, Habitus, dorsal view; B, Cephalon; C, Antennule; D, Antenna; E, Pleotelson and uropod; F, Penes. Scale bars: A, B = 0.5 mm, C−E = 0.2 mm, F = 0.1 mm. (cited from Kim & Yoon, 2019b) ··· 95 Fig. 37. Elaphognathia kikuchii, male. A, Mandible, dorsolateral view; B, Maxilliped; C, Distal end of pylopod, D, Pylopod. Scale bars: A, C = 0.2 mm, B, D = 0.5 mm. (cited from Kim & Yoon, 2019b) ··· 97 Fig. 38. Elaphognathia kikuchii, male. A, Pereopod 2; B, Pereopod 3; C, Pereopod 4; D, Pereopod 5; E, Pereopod 6. Scale bar: A−E = 0.2 mm. (cited from Kim & Yoon, 2019b) ··· 99 Fig. 39. Elaphognathia kikuchii, male. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5. Scale bar: A−E = 0.5 mm. (cited from Kim & Yoon, 2019b) ··· 101 Fig. 40. Elaphognathia kikuchii, female. A, Habitus, dorsal view; B, Cephalon; C, Pleotelson; D, Antennule; E, Antenna; F, Maxilliped; G, Pylopod; H, Pereopod 2; I, Pleopod 2. Scale bars: A
vii
= 0.5 mm, B−G = 0.2 mm, H, I = 0.1 mm. ··· 103 Fig. 41. Elaphognathia monodi, male. A, Habitus, dorsal view; B, Cephalon; C, Pleotelson and uropod; D, Antennule; E, Antenna; F, Penes. Scale bars: A, B = 0.5 mm, C−E = 0.2 mm, F = 0.1 mm. (cited from Kim & Yoon, 2019b) ··· 105 Fig. 42. Elaphognathia monodi, male. A, Mandible, dorsal view; B, Maxilliped; C, Distal end of pylopod; D, Pylopod. Scale bars: A, B, D = 0.2 mm, C = 0.1 mm. (cited from Kim & Yoon, 2019b) ··· 107 Fig. 43. Elaphognathia monodi, male. A, Pereopod 2; B, Pereopod 3; C, Pereopod 4; D, Pereopod 5;
E, Pereopod 6. Scale bar: A−E = 0.2 mm. (cited from Kim & Yoon, 2019b) ··· 109 Fig. 44. Elaphognathia monodi, male. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5. Scale bar: A−E = 0.5 mm. (cited from Kim & Yoon, 2019b)··· 111 Fig. 45. Gnathia n. sp., holotype, male. A, Habitus, dorsal view; B, Cephalon, dorsal view; C, Cephalon, ventral view; D, Cephalon, lateral view; E, Mandible, lateral view; F, Antennule; G, Antenna; H, Maxilliped; I, Pylopod. Scale bars: A = 1 mm, B−D = 0.5 mm, E−I = 0.2 mm.
··· 113 Fig. 46. Gnathia n. sp., holotype, male. A, Pereopod 2; B, Pereopod 3; C, Pereopod 4, D, Pereopod 5; E, Pereopod 6. Scale bar: A−E = 0.2 mm. ··· 115 Fig. 47. Gnathia n. sp., holotype, male. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F. Pleotelson and uropod. Scale bar: A−F = 0.5 mm. ··· 117 Fig. 48. Bopyrinae n. gen. & n. sp., holotype, female. A, Habitus, dorsal view; B, Habitus, ventral view; C, Maxilliped and barbular; D, Left oostegite 1; E, Right oostegite 1; F, Pereopod 1; G, Pereopod 2; H, Pereopod 7. Scale bars: A, B = 2 mm, C−E = 0.5 mm, F−H = 0.25 mm. ··· 122 Fig. 49. Parabopyrella n. sp., holotype, female (A−F) and male (G−K). A, Habitus, dorsal view; B, Habitus, ventral view; C, Maxilliped and barbular; D, Right oostegite 1; E, Left oostegite 1; F, Pereopod 7; G, Habitus, dorsal view; H, Habitus, ventral view; I, Antennule; J, Antenna; K, Pereopod 7. Scale bars: A, B = 4 mm, D, E = 1 mm, C, F−H = 0.5 mm; I−K = 0.05 mm.
··· 124
viii
Fig. 50. Limnoria n. sp., holotype, female. A, Habitus, dorsal view; B, Habitus, lateral view; C, Pleotelson, dorsolateral view; D, Pleotelson, dorsal view; E, Antennule; F, Antenna; G, Right mandible; H, Left mandible; I, Maxillule; J, Maxilla; K, Maxilliped. Scale bars: A−D = 0.5 mm, E−K = 0.1 mm. ··· 128 Fig. 51. Limnoria n. sp., holotype, female. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 0.2 mm. ··· 130 Fig. 52. Limnoria n. sp., holotype, female. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4;
E, Pleopod 5; F, Uropod. Scale bar: A−F = 0.2 mm. ··· 132 Fig. 53. Idarcturus n. sp., holotype, female. A, Habitus, dorsal view; B, Habitus, lateral view; C, Oostegite 4. Paratype, male: D, Habitus, dorsal view; E, Habitus, lateral view; F, Pleopod 2.
Scale bars: A, B, D, E = 1 mm, C, F = 0.2 mm. ··· 150 Fig. 54. Idarcturus n. sp., holotype, female. A, Antennule; B, Antenna; C, Detail of peduncular articles of antenna; D, Flagellum of antenna; E, Left mandible; F, Right mandible; G, Maxillule;
H, Maxilla; I, Maxilliped. Scale bars: A, B, D = 0.2 mm, E−I = 0.1 mm, C = 0.02 mm. ···· 152 Fig. 55. Idarcturus n. sp., holotype, female. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bars: A−G = 0.1 mm. ···· 154 Fig. 56. Idarcturus n. sp., holotype, female. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F, Uropod. Scale bars: A−F = 0.2 mm. ··· 156 Fig. 57. Neastacilla n. sp., holotype, male. A, Habitus, dorsal view; B, Habitus, lateral view; E, Penes.
Paratype, female: C, Habitus, dorsal view; D, Habitus, lateral view. Scale bars: A−D = 1 mm, E = 0.1 mm. ··· 159 Fig. 58. Neastacilla n. sp., holotype, male. A, Antenna; B, Distal end of antenna; C, Antennule; D, Maxillule, E, Maxilla; F, Right mandible; G, Left mandible; H, Maxilliped. Scale bars: A, B = 0.5 mm, C−H = 0.2 mm. ··· 161 Fig. 59. Neastacilla n. sp., holotype, male. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 0.5 mm. ··· 163 Fig. 60. Neastacilla n. sp., holotype, male. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4;
ix
E, Pleopod 5; F, Uropod. Scale bars: A−F = 0.5 mm. ··· 164 Fig. 61. Symmius caudatus, female. A, Habitus, dorsal view; B, Habitus, lateral view; C, Antennule;
D, Antenna. Scale bars: A−D = 1 mm. (cited from Kim & Yoon, 2018b) ··· 166 Fig. 62. Symmius caudatus, female. A, Left mandible; B, Right mandible; C, Left maxillule; D, Left maxilla; E, Right maxilla; F, Maxilliped. Scale bar: A−F = 0.5 mm. (cited from Kim & Yoon, 2018b) ··· 168 Fig. 63. Symmius caudatus, female. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4;
E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 1 mm. (cited from Kim & Yoon, 2018b) ··· 169 Fig. 64. Symmius caudatus, female. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F, Uropod. Scale bars: A−F = 1 mm. (cited from Kim & Yoon, 2018b) ··· 171 Fig. 65. Symmius caudatus, male. A, Habitus, dorsal view; B, Habitus, lateral view; C, Antennule;
D, Pereopod 1; E, Pereopod 7; F, Pleopod 2; G, Penes. Scale bars: A = 1 mm, B−F = 0.5 mm, G = 0.25 mm. (cited from Kim & Yoon, 2018b) ··· 173 Fig. 66. Symmius caudatus, Juvenile male. A, Habitus, dorsal view; B, Habitus, lateral view; C, Pleopod 2; D, Pereopod 7. Scale bars: A, B = 1 mm, C, D = 0.25 mm. (cited from Kim & Yoon, 2018b) ··· 174 Fig. 67. Cleantiella n. sp., holotype, male. A, Habitus, dorsal view; B, Pleon, ventral view; C, Penes;
D, Antennule; E, Antenna; F, Left mandible; G, Right mandible; H, Maxillule; I, Maxilla; J, Maxilliped. Scale bars: A = 2 mm, B, E = 0.5 mm, C, D, F−J = 0.2 mm. ··· 179 Fig. 68. Cleantiella n. sp., holotype, male. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 1 mm. ··· 181 Fig. 69. Cleantiella n. sp., holotype, male. A, Pleopod 1; B, Pleopod 2; C, Distal end of appendix masculina; D, Pleopod 3; E, Pleopod 4; F, Pleopod 5; G, Uropod. Scale bars: A, B, D−G = 0.5 mm, C = 0.1 mm. ··· 183 Fig. 70. Cleantiella n. sp., female (A, B) and juvenile male (C−F). A, Habitus, ovigerous female; B, Habitus, not ovigerous female; C, Habitus, dorsal view; D, Habitus, lateral view; E, Antennule;
x
F, Pleopod 2. Scale bars: A, B = 2 mm, C, D, F = 1 mm, E = 0.1 mm. ··· 185 Fig. 71. Pentias n. sp., holotype, male. A, Habitus, dorsal view; B, Habitus, lateral view; C, Cephalon;
D, Pereonite 7 to pleotelson, ventral view. Scale bars: A, B = 5 mm, C, D = 1 mm. ··· 191 Fig. 72. Pentias n. sp., holotype, male. A, Antennule; B, Antenna; C, Distal end of antenna; D, Left mandible; E, Right mandible; F, Maxillule; G, Maxilla; H, Maxilliped. Scale bars: A, C = 0.1 mm, B = 0.5 mm, D−H = 0.2 mm. ··· 193 Fig. 73. Pentias n. sp., holotype, male. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 0.5 mm. ··· 195 Fig. 74. Pentias n. sp., holotype, male. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F, Uropod. Scale bars: A−F = 1 mm. ··· 197 Fig. 75. Janiralata sagamiensis, male. A, Habitus; B, Lateral margin of habitus; C, Pleotelson; D, Antennule; E, Antenna; F, Left mandible; G, Right mandible; H, Maxillule; I, Maxilla; J, Maxilliped. Scale bars: A = 1 mm, B = 0.5 mm, C−J = 0.1 mm. ··· 206 Fig. 76. Janiralata sagamiensis, male. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 0.5 mm. ··· 208 Fig. 77. Janiralata sagamiensis, male (A−F) and female (G). A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F, Uropod; G, Pleopod 1. Scale bars: A−G = 0.2 mm. ··· 210 Fig. 78. Joeropsis n. sp. 1, holotype, male. A, Habitus, dorsal view; C, Pseudorostrum; D, Antennule;
E, Antenna. Paratype, female: B, Habitus, dorsal view. Scale bars: A, B = 0.5 mm, C−E = 0.1 mm. ··· 213 Fig. 79. Joeropsis n. sp. 1, holotype, male. A, Left mandible; B, Right mandible; C, Maxillule; D, Maxilla; F, Left maxilliped; G, Distal end of left maxillipedal endite; H, Right maxilliped; I, Distal end of right maxillipedal endite. Scale bars: A, B, F, H = 0.2 mm, D, E = 0.1 mm, C, G, I = 0.2 mm. ··· 215 Fig. 80. Joeropsis n. sp. 1, holotype, male. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 0.5 mm. ··· 217 Fig. 81. Joeropsis n. sp. 1, holotype, male. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4;
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E, Pleopod 5; G, Uropod. Paratype, female: F, Pleopod 1. Scale bars: A−F = 0.1 mm, G = 0.5 mm. ··· 219 Fig. 82. Joeropsis n. sp. 2, holotype, male: A, Habitus, dorsal view; C, Lateral margin of pleotelson;
D, Pseudorostrum; E, Antennule; F, Antenna. Paratype, female: B, Habitus, dorsal view. Scale bars: A−F = 0.1 mm. ··· 221 Fig. 83. Joeropsis n. sp. 2, holotype, male. A, Left mandible; B, Right mandible; C, Maxillule; D, Maxilla; E, Left maxilliped; F, Distal end of maxillipedal endites. Scale bars: A−F = 0.1 mm. ··· 223 Fig. 84. Joeropsis n. sp. 2, holotype, male. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar: A−G = 0.1 mm. ··· 225 Fig. 85. Joeropsis n. sp. 2, holotype, male. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4;
E, Pleopod 5; F, Uropod. Paratype, female: G, Pleopod 1. Scale bars: A−E, G = 0.1 mm, F = 0.02 mm. ··· 227 Fig. 86. Stenetrium sp., female. A, Habitus; B, Lateral margin of pereonites; C, Pleotelson; D, Antennule; E, Antenna. Scale bars: A, B = 1 mm, C, E = 0.5 mm, D = 0.2 mm. ··· 229 Fig. 87. Stenetrium sp., female. A, Left mandible; B, Right mandible; C, Maxillule, D, Maxilla; E, Maxilliped; F, Distal end of endite; G, Spines. Scale bars: A−E = 0.2 mm, F = 0.1 mm, G = 0.01 mm. ··· 231 Fig. 88. Stenetrium sp., female. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bars: A, C−H = 1 mm, B = 0.2 mm. ··· 233 Fig. 89. Stenetrium sp., male (A−E, G) and female (F). A, Pleopod 1; B, Pleopod 2; C, Pleopod 3;
D, Pleopod 4; E, Pleopod 5; F, Pleopod 2; G, Uropod. Scale bar: A−G = 0.2 mm. ··· 235 Fig. 90. Cladograms based on unweighted data (length = 134, consistency index = 0.55, retention index = 0.87). A, Strict consensus of 215 trees; B, Majority-rule consensus of 215 trees.
Percentage (%) of occurrence in the 215 equally most parsimonious trees is shown at nodes (B).
··· 272 Fig. 91. Cladogram by majority-rule consensus of 36 trees (length = 67.56, consistency index = 0.78,
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retention index = 0.94), based on reweighted data. Clade number, percentage (%) of occurrence in the 36 equally most parsimonious trees, and posterior probabilities for the Bayesian inference are shown at each node in sequence. A dash means the support below 50% in the Bayesian inference. ··· 273 Fig. 92. Cladogram by bootstrap 50% majority-rule consensus of 36 trees, based on reweighted data.
The maximum parsimony algorithm was used to depict tree. Bootstrap values for the maximum parsimony are shown at nodes. ··· 274 Fig. 93. Early cladograms of isopod taxa. A, Cladogram of Wägele (1989) combined by Brandt &
Poore (2003: fig. 7); B, Cladogram postulated by Brusca & Willson (1991: fig. 14); C, Cladogram by Brandt & Poore (2003). ··· 281 Fig. 94. An evolutionary scenario for the parasitic isopods based on the results of the majority-rule consensus of 36 trees (except for gnathiid isopods). The arrows mean the direction of evolution.
Illustrations of Corallanidae and Aegidae were referred to Song & Min (2018) and National Institute of Biological Resources (2012). ··· 282
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LIST OF TABLES
Table 1. Comparison of known species of the genus Cleantiella.··· 186 Table 2. Comparison of known species of the genus Pentias. ··· 198 Table 3. Species used in the morphological analysis. ··· 252 Table 4. Characters and character states used in the cladistics analysis of the Isopoda (referred on Brandt & Poore, 2003). ··· 260 Table 5. Character matrix of 54 isopod taxa and 59 characters used in the cladistic analysis. ···· 263 Table 6. Synapomorphies of each clade and taxon in the 50% majority-rule consensus of 36 trees, based on reweighted data. ··· 275
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국문초록
김 성 훈
지도교수: 윤 성 명 생명과학과
조선대학교 대학원
등각류는 암컷의 복부에 위치한 육아낭(marsupium)으로 자신의 유생을 기르는 낭하상목(Peracarida)에 속하며, 육상, 담수, 기수 및 해양의 다양한 서식처에서 발견 된다. 국내에서는 약 120여종의 등각류가 다양한 환경에서 보고되었으며, 그 중 약 80여종이 해양성으로 알려져 있다.
본 연구는 등각류(isopods)에 대한 계통분류학적 연구로서, 두 부분으로 구성되 었다: Part 1, 한국 해양성 등각류의 형태학적 연구; Part 2, 한국산 재료를 이용한 등 각류의 계통분류학적 연구. Part 1의 등각류의 형태학적인 연구는 2014년부터 2019 년까지 해양의 조간대와 조하대를 포함하는 남한 152개의 정점에서 채집된 표본을 대상으로 수행되었다. 그 결과, 4아목, 17과, 44속에 포함되는 72종의 등각류를 확인하 였고, 그 중에는 15종의 신종 후보와 8종의 국내미기록종 후보(Amakusanthura n. sp.;
Apanthura n. sp.; Caenanthura engimatica (Kensley & Reid, 1984); Kupellonura n. sp. 1;
Kupellonura n. sp. 2; Eurydice longiantennata Nunomura & Ikehara, 1985; Metacirolana japonica (Hansen, 1890); Nerocila sp.; Elaphognathia kikuchii (Nunomura, 1992); Elaphognathia monodi (Gurjanova, 1936); Gnathia n. sp; Bopyrinae n. gen. & n. sp.; Parabopyrella n. sp.; Limnoria n. sp.;
Idarcturus n. sp.; Neastacilla n. sp.; Symmius caudatus Richardson, 1904; Cleantiella n. sp.; Pentias n. sp.; Janiralata sagamiensis Shimomura, 2006; Joeropsis n. sp. 1; Joeropsis n. sp. 2; Stenetrium sp.)가 포함되어 있다. 본 연구에서는 이들 23종의 신종후보 및 국내미기록종에 대한 자세한 기재와 도판이 작성되었으며, 기수역을 포함한 한국 연안에서 채집된 등각류에
xv 대한 검색표를 작성하였다.
Part 2에서는 30과 54속에 포함되는 54개 종들에 대한 59개 형태적 형질들을 분석하여 이를 바탕으로 한 등각목에 대한 계통수를 작성하였다. 주요 분류군 중에서 본 연구에서는 채집하지 못한 Calabozoa, Coxicerberus, Phreatoicus, Phoratopus, Tachaea, 및 Tainisopus의 6개 속의 종들은 문헌을 바탕으로 형태적 형질을 확인하였으며, 이들 을 제외한 모든 종들은 직접 관찰하여 형질을 확인하였다. 형질에 가중치를 부여하지 않은 첫 번째 분석에서는 215개의 계통수(steps=134, CI=0.55, RI=0.871)가 확인되었고, 가중치를 부여한 두 번째 분석에서는 36개의 계통수(steps=67.56, CI=0.78, RI=0.94)가 얻 어졌다. 본 연구의 결과 얻어진 계통수는 이전 연구의 많은 주장들 가운데 다음과 같 은 결론을 지지했다: (1) Microcerberidea는 Asellota와 자매군(sister group)을 형성하며, 가 장 원시적인 등각류라고 알려진 Phreatoicidea와 함께 계통수의 가장 아래에 위치한다;
(2) Tainisopids는 분류계급적으로 아목으로 분류되었지만 이를 뒷받침할 강력한 증거는 확인되지 않는다; (3) 유변아목(Valvifera)과 잔벌레아목(Sphaeromatidea)의 자매군 관계가 강력하게 지지된다; (4) Calabozoidea는 계통수에서 쥐며느리아목(Oniscidea)의 아래에 위 치하면서 이 아목과 자매군을 형성한다; (5) Phoratopidea와 부삽꼬리벌레아목 (Limnoriidea)은 유변아목이나 잔벌레아목(Sphaeromatidea)보다 부채벌레아목(Cymothoida) 에 가깝게 위치한다; (6) 모래무지벌레과(Cirolanaidae)는 부채벌레아목 내에서 가장 아 래에 위치한다; (7) Epicaridea는 갈고리벌레과(Cymothoidae)와 자매군을 형성하며, 부채 벌래아목에 속하는 하나의 과로 분류될 수 있다.
１
Part 1. Mophological Study of the Marine Isopods
(Crustacea, Peracarida) from Korea
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1. Introduction
The isopods are one of the peracarids that brood their offsprings in marsupium under the pereon (Wetzer et al., 1997; Brusca et al., 2001; Poore & Bruce, 2012). The peracarids consisted of 12 orders:
Amphipoda Latreille, 1816; Isopoda Latreille, 1817; Lophogastrida Boas, 1883; Mysida Boas, 1883;
Cumacea Krøyer, 1846; Tanaidacea Dana, 1849; Ingolfiellida Hansen, 1903; Thermosbaenacea Monod, 1927; Spelaeogriphacea Gordon, 1957; Stygiomysida Tchindonova, 1981; Mictacea Bowman, Garnet, Hessler, Iliffe & Sanders, 1985; and Bochusacea Gutu & Iliffe, 1998 (Brusca et al., 2001; Martin & Davis, 2001; WoRMS, 2019). Among them, the isopods can be distinguished from other peracarids in the following features: (1) body is dorsoventrally flattened and divided into three parts: a cephalon, pereon, and pleon; (2) coxae are small and simple or almost fused to pereonites extending laterally or ventrally; (3) antennule and antenna are uniramous; (4) seven pairs of pereopods are uniramous; (5) six pairs pleonal appendages are biramous (Kensley, 1978; Brusca &
Wilson, 1991; Brusca et al., 2001; Kazmi et al., 2002; Poore & Bruce, 2012). Currently, the order Isopoda is divided into the following eleven suborders: Asellota Latreille, 1802; Oniscidea Latreille, 1802; Valvifera G.O. Sars, 1883; Phreatoicidea Stebbing, 1893; Microcerberidea Lang, 1961;
Calabozoidea Nan Lieshout, 1983; Cymothoida Wägele, 1989; Sphaeromatidea Wägele, 1989;
Limnoriidea Brandt & Poore in Poore, 2002; Phoratopidea Brandt & Poore, 2003; and Taninisopidea Brandt & Poore, 2003 (Brandt & Poore, 2003; Poore & Bruce, 2012).
The isopods live in various environments such as terrestrial, freshwaters, estuarine, and marine habitats conducting important roles as intermediate links in the food web. In the sea, they are predominant together with tanaids, bivalves, and polychaetes and affect the environment and organisms including humans in various ways (e.g. some cirolanids attack fish or humans; Limnoria damage to wooden boats and structures boring into the wood) (Poore & Bruce, 2012). About 10,000 species are recorded worldwide. Among them, over 6,000 species are known from estuarine or marine waters (Brusca et al., 2001; Brandt & Poore, 2003; Poore & Bruce, 2012). Despite their species diversity, in Korean waters, only about 120 species have been reported from various environments including terrestrial, freshwaters, brackish, and marine ones. Among them, about 80
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species are reported from estuarine or marine habitats (National Institute of Biological Resources, 2019).
Early studies of the isopods fauna in Korea were mainly carried by foreign researchers (Richardson, 1909; Iwasa, 1965). After to this, a lot of taxonomic studies had been performed by native researchers during 1950s to 1990s (Kim & Kown 1985; 1988a, b; Kwon & Kim, 1986; 1987;
1992; Kwon; 1986; 1988; 1990a, b; 1995; 1998; Jang, 1991; Jang & Kwon, 1990; 1993). While a vacancy has existed (Malyutina & Ushakova, 2001a, b), the study of Korean isopods are currently presented at a high frequency by several authors (Song et al, 2014; 2017; Song & Min, 2015a, b, c;
2016a, b; 2017a, b, c; 2018a, b; Hong et al., 2015).
A taxonomic study was conducted to elucidate the marine isopod fauna of Korea. The materials were collected from various marine habitats at 152 localities including intertidal and subtidal zones in Korean waters during the periods from 2014 to 2019. As a result, 72 species belonging to four suborders, 17 families, and 44 genera were collected. Among them, 23 species were identified as the species new to science or new to Korean fauna. The list of them is as follows: Amakusanthura n. sp.;
Apanthura n. sp.; Caenanthura engimatica (Kensley & Reid, 1984); Kupellonura n. sp. 1;
Kupellonura n. sp. 2; Eurydice longiantennata Nunomura & Ikehara, 1985; Metacirolana japonica (Hansen, 1890); Nerocila sp.; Elaphognathia kikuchii (Nunomura, 1992); Elaphognathia monodi (Gurjanova, 1936); Gnathia n. sp.; Bopyrinae n. gen. & n. sp.; Parabopyrella n. sp.; Limnoria n. sp.;
Idarcturus n. sp.; Neastacilla n. sp. Symmius caudatus Richardson, 1904; Cleantiella n. sp.; Pentias n. sp.; Janiralata sagamiensis Shimomura, 2006; Joeropsis n. sp. 1; Joeropsis n. sp. 2; Stenetrium sp. These 23 species were described and illustrated in detail, with keys to all taxa of Korean isopods in this study.
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2. Materials and methods
During an expedition from 2014 to 2019, the materials of the present study were collected at 152 localities including intertidal to subtidal zones in Korean waters (Fig. 1). Among them, the intertidal materials were taken by a sieve with a mesh size of 1 mm or hand. The subtidal materials were sampled by using a van Veen or smith-Mclntyre grab, light trap, and SCUBA diving ranging from 1 m to 60 m depth. The collected materials were immediately fixed in 95% ethanol and then transferred to the laboratory. The transferred materials were sorted and dissected under the stereomicroscope (Nikon SMZ 1500, Japan) and then observed using a bright-field microscope (Olympus, BX 50, Japan). The dissection was carried out under the glycerin. After finishing observation of the whole specimens and each appendage, the examined materials were transferred to Canada balsam to make permanent slides or small glass vials filled with 95% ethanol to preserve the dissected materials. Measurements and drawings of the whole specimens were performed with the aid of a drawing tube and camera lucida. The drawings were scanned, inked, and arranged digitally using Adobe Illustrator CS6. The drawing methods followed by Coleman (2003, 2009). All examined materials were deposited at the National Institute of Biological Resource (NIBR) and Chosun University in Korea.
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Fig. 1. Localities of the present study (the list is provided in Appendix 1).
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3. Morphology and taxonomic characters
The classification scheme of Isopoda was based on Brandt & Poore (2003). The terminology for detailed morphology and taxonomic characters followed the representative works by several previous authors (Kensley, 1978; Brusca & Iverson, 1985; Brandt & Poore, 2003).
3.1. General morphology 1) Body
Order Isopoda lacks carapace covering the pereonites. As a result, the pereonites are clearly exposed. The body is generally depressed dorsoventrally, occasionally cylindrical (e.g. anthurideans) or laterally flattened (e.g. Phreatoicidea) and is divided into three parts: the cephalon (head), pereon (thorax), and pleon (abdomen). The cephalon and first thorax are always fused to each other and the first thoracopods modified into maxillipeds. So, the cephalothorax is referred to as the cephalon due to pereonite 2 means pereonite 1 conventionally.
2) Cephalon (head)
The cephalon bears the sets of paired appendages: antennule, antenna, mandible, maxillule, maxilla, and maxilliped. The antennae are usually uniramous except for a few genera that have vestigial exopod represented by the antennal scale (e.g. asellotans). Eyes are compound eyes having numerous ommatidia, sometimes composed of ocellus in the family Trichoniscidae (Oniscidea). In some isopods, the anterior margin of cephalon is expanding anteriorly forming the rostrum (frontal process).
(1) Antennule: the first pair antennae (antennule) are usually composed of three peduncular articles and a flagellum. The number of flagellar articles are various depends on the taxa.
(2) Antenna: the second antennae (antenna) normally consists of five peduncular and numerous flagellar articles. The number of flagellar articles of the antenna depends on the taxa.
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(3) Frontal lamina is located between both antennae. The structure is considered as the homology of the epistome commonly formed to all groups of crustaceans.
(4) Labrum is positioned on the posterior region of the frontal lamina. The structure consists of two parts: the anterior part referred to clypeus and the posterior parts called as labrum.
(5) Mandible is composed of 1−3-articled palp, sclerotized incisor process, articulated lacinia mobilis, and a molar process, occasionally a spine row positioned between the incisor and molar process. The structure of mandible is variously modified depend on the taxa.
(6) Maxillule consists of two lobes, inner and outer lobes. The inner lobe functions normally sensory and the outer lobe conducts as masticatory or biting.
(7) Maxilla is composed of two lobes, inner and outer lobes. The outer lobe is usually bifurcated partly or entirely. The bifuracated lob is called as mesial and outer lobes.
(8) Maxilliped is primitively derived from pereopod since the first thorax are fused to cephalon.
The coxa bears a normally separated lobe, inner and outer lobes. The outer lobe is called as epipod.
The inner lobe is composed of the modified articles. The basis is enlarged and has an endite and coupling hook. The other articles are referred to as palp. The palp is variously fused depending on the taxa.
2) Pereon (thorax)
The pereon is composed of pereonites which are not fused to the cephalon. It is normally 7- segmented and each pereonite bearing a pair of uniramous appendages. Unusually, pereonite 1 of gnathiids is fused to cephalon, so that pereopod 1 modified as pylopod.
(1) Pereopod is usually 7-articled: coxa, basis, ischium, merus, carpus, propodus, and dactylus.
The coxa is referred to as the coxal plate. It is expanded and fused partly or completely to the lateral margin of each pereonites. The coxal plates only observed in both peracarid taxa, isopods and amphipods, are diversely fused to pereonites depending on the taxa. In the female, several coxae expand mesially, forming thin plates on the ventral surface. The expanded coxae are referred to as oostegites. They form the brood pouch or marsupium. The pereopods have various forms, e.g.
８
ambulatory, natatory, subchelate, or prehensile. The cheliped form has not occurred in isopods. The ambulatory form is simple and used for walking. The natatory form has a flattened basis for swim.
The subchelate and prehensile forms are strongly recurved and used for grasping. The two later forms can be distinguished by the relative length of dactlyus to porpous. The dactylus of prehensile form is similar to or longer than propodus, whereas dactylus of subchelate form shorter is than propodus.
3) Pleon (abdomen)
The pleon consists of variously fused segments. Each segment has a pair of biramous appendages. Generally, the terminal pleonite is fused to telson and it is called pleotelson. The anus is positioned on the subterminal region of pleotelson. Occasionally, the anus is located on the dorsal side of pleotelson in some taxa (e.g. anthuridean) and it is called to statocysts.
(1) Pleopod is laminated biramous and it functions as gills and swimming appendages. Depend on the taxa, the rami have several plumose setae along the margin. In the male, pleopod 2 has stylet on the inner margin of endopod. In some taxa (e.g. anthuroidean and valviferan), the pleopods are covered by uropods or other pleopods.
(2) Uropod is composed of protopod (sympod or peduncle), exopod, and endopod. The uropod is used in the higher classification of Isopoda because they have diverse morphologies depend on the higher taxa. Several taxa (e.g. Asellota and Limnoriidea) have styliform uropods whereas some taxa (e.g. Cymothoida and Sphaeromatidea) have broad and flat uropods. In the Valvifera, the uropods are positioned on the ventral side, covering the pleopods.
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Fig. 2. General morphology of body. A, Habitus, dorsal view (left, Metacirolana; right, Janiralata);
B, Habitus, lateral view (Metacirolana).
１０
Fig. 3. General morphologies of cephalon and cephalic appendages. A, Antennule (Chaetiliidae); B, Antenna (Janiridae); C, Ventral view of cephalon (Cirolanidae); D, Mandible (Idoteidae); E, Mandible (Cirolanidae); F, Maxillule (Cirolanidae); G, Maxilla (Cirolanidae); H, Maxilliped (Idoteidae).
１１
Fig. 4. Morphologies of pereopods. A, Prehensile pereopod (Cymothoidae); B, Subchelate pereopod (Anthuridae); C, Ambulatory pereopod (Cirolanidae); D, Natatory pereopod (Cirolanidae); E Pylopod (Gnathiidae).
１２
Fig. 5. Morphologies of pleonal appendages. A, Pleopod 1 (Anthuridae); B, Pleopod 2 of male (Idoteidae); C, Uropod (Janiridae); D, Uropod (Cirolanidae); E, Uropod (Idoteidae); F, Uropod (Anthuridae); G, Pleotelson (Anthuridae).
１３
4. Systematic accounts
4.1. List of species
Order Isopoda Latreille, 1817 등각목
Suborder Cymothoida Wägele, 1989 부채벌레아목 Superfamily Anthuroidea Leach, 1914 마디벌레상과 Family Anthuridae Leach, 1814 마디벌레과 Genus Amakusanthura Nunomura, 1977
**1. Amakusanthura n. sp.
Genus Apanthura Stebbing, 1900 길쭉마디벌레속
**2. Apanthura n. sp.
3. Apanthura koreaensis Song & Ming, 2015 길쭉마디벌레 Genus Caenanthura Kensley, 1978 무목마디벌레속
*4. Caenanthura engimatica (Kensley & Reid, 1984) 등홈무목마디벌레 Genus Cyathura Norman & Stebbing, 1886
5. Cyathura higoensis Nunomura, 1977 모래마디벌레 Genus Mesanthura Barnard, 1914
6. Mesanthura nigrodorsalis Nunomura, 1977 검은마디벌레 Family Hyssuridae Wägele, 1981
Genus Kupellonura Barnard, 1925
**7. Kupellonura n. sp. 1
**8. Kupellonura n. sp. 2
Family Paranthuridae Menzies & Glynn, 1968 오목꼬리마디벌레과 Genus Colanthura Richardson, 1902 배마디벌레속
9. Colanthura pigmentata Kensley, 1980 깜장배마디벌레 Genus Paranthura Bate & Westwood, 1866
１４ 10. Paranthura japonica Richardson, 1909 큰마디벌레 Superfamily Cymothoidea Wägele, 1989 부채벌레상과 Family Aegidae White, 1850 부채벌레과
Genus Rocinela Leach, 1818 부채벌레속
11. Rocinela maculata Schioedte & Meinert, 1879 네점박이부채벌레 Family Cirolanidae Dana, 1852 모래무지벌레과
Genus Cirolana Leach, 1818 가시꼬리모래무지벌레속 12. Cirolana koreana Kwon, 1988 얼룩모래무지벌레
13. Cirolana toyamaensis Nunomura, 1982 울퉁모래무지벌레 Genus Eurydice Leach, 1815 꺽인더듬이모래무지벌레속
*14. Eurydice longiantennata Nunomura & Ikehara, 1985 긴꺽인더듬이모래무지벌레 Genus Excirolana Richardson, 1912 모래무지벌레속
15. Excirolana chiltoni Richardson, 1912 모래무지벌레
Genus Metacirolana Kussakin, 1979 톱니꼬리모래무지벌레속
*16. Metacirolana japonica (Hansen, 1890) 긴허리톱니꼬리모래무지벌레 Genus Natatolana Bruce, 1981 헤엄다리모래무지벌레속
17. Natatolana japonensis (Richardson, 1904) 왜모래무지벌레 Family Cymothoidae Leach, 1814 갈고리벌레과
Genus Mothocya Hope, 1851 아감벌레속
18. Mothocya melanosticta (Schioedte & Meinert, 1884) 학공치아감벌레 Genus Nerocila Leach, 1818
*19. Nerocila sp.
Family Gnathiidae Leach, 1814 큰턱벌레과
Genus Elaphognathia Monod, 1926 오목머리큰턱벌레속
１５
*20. Elaphognathia kikuchii (Nunomura, 1992) 수염오목머리큰털벌레
*21. Elaphognathia monodi (Gurjanova, 1936) 네돌기오목머리큰턱벌레
22. Elaphognathia sugashimaensis (Nunomura, 1981) 스가시마오목머리큰턱벌레 Genus Gnathia Leach, 1814 큰턱벌레속
**23. Gnathia n. sp.
23. Gnathia koreana Song & Min, 2018 한국큰턱벌레 Infraorder Epicaridea Latreille, 1831
Superfamily Bopyroidea Rafinesque, 1815
Family Bopyridae Rafinesque, 1815 십각기생과 Subfamily Athelginae Codreanu & Codreanu, 1956
Genus Parathelges Bonnier, 1900
25. Parathelges enoshimensis Shiino, 1950 참집게배붙이 Subfamily Bopyrinae Rafinesque, 1815
Genus Bopyrinae n. gen.
**26. Bopyrinae n. gen. & n. sp.
Genus Parabopyrella Markham, 1985
**27. Parabopyrella n. sp.
Suborder Limnoriidea Brandt & Poore in Poore, 2002 부삽꼬리벌레아목 Superfamily Limnorioidea Whith, 1850 부삽꼬리벌레상과
Family Limnoriidae Leach, 1814 부삽꼬리벌레과 Genus Limnoria Leach, 1814 부삽꼬리벌레속
**28. Limnoria n. sp.
29. Limnoria rhombipunctata Yoshino, Watabe & Ohsawa, 2017 동해부삽꼬리벌레 30. Limnoria segnoides Menzies, 1957 작은혹부삽꼬리벌레
Suborder Sphaeromatidea Wägele, 1989 잔벌레아목 Superfamily Sphaeromatoidea Latreille, 1825 잔벌레상과
１６ Family Sphaeromatidae Latreille, 1825 잔벌레과
Genus Chitonosphaera Kussakin & Malyutina, 1993 납작잔벌레속 31. Chitonosphaera lata (Nishimura, 1968) 납작잔벌레
Genus Cymodoce Leach, 1814 혹잔벌레속
32. Cymodoce acuta Richardson, 1904 두드러기잔벌레 33. Cymodoce japonica Richardson, 1907 두혹잔벌레 Genus Dynamenella Hansen, 1905 홈잔벌레속
34. Dynamenella nipponica (Nishimura, 1969) 두드러기홈잔벌레 Genus Dynoides Barnard, 1914 뿔잔벌레속
35. Dynoides brevicornis Barnard, 1914 뭉퉁뿔잔벌레 36. Dynoides brevispina Bruce, 1980 넓적뿔잔벌레 37. Dynoides dentisinus Shen, 1929 가시홈뿔잔벌레 38. Dynoides longisinus Kwon, 1990 긴홈뿔잔벌레
39. Dynoides spinipodus Kwon & Kim, 1986 가시다리뿔잔벌레 Genus Gnorimosphaeroma Menzies, 1954 잔벌레속
40. Gnorimosphaeroma chejuense Kim & Kwon, 1988 제주잔벌레 41. Gnorimosphaeroma hoestlandti Kim & Kwon, 1985 꼬마갯가잔벌레 42. Gnorimosphaeroma naktongense Kwon & Kim, 1987 낙동잔벌레 43. Gnorimosphaeroma ovatum (Gurjanova, 1933) 개펄잔벌레 44. Gnorimosphaeroma rayi Hoestlandt, 1969 갯가잔벌레 45. Gnorimosphaeroma noblei Menzies, 1954 길쭉잔벌레 Genus Holotelson Richardson, 1909 세혹잔벌레속 46. Holotelson tuberculatus Richardson, 1909 세혹잔벌레 Genus Leptosphaeroma Hilgendorf, 1885 납작잔벌레붙이속
１７
47. Leptosphaeroma gotschei Hilgendorf, 1885 납작잔벌레붙이 Genus Sphaeroma Bosc, 1801 큰잔벌레속
48. Sphaeroma sieboldii Dollfus, 1889 일곱니잔벌레 Suborder Valvifera Sars, 1882 유변아목
Family Arcturidae Dana, 1849 기린벌레과 Genus Idarcturus Barnard, 1914
**49. Idarcturus n. sp.
Genus Neastacilla Tattersall, 1921
**50. Neastacilla n. sp.
Family Chaetiliidae Dana, 1849 두엽벌레과 Genus Symnius Richardson, 1904 창벌레속
*51. Symmius caudatus Richardson, 1904 창벌레
52. Symnius yamaguchiensis Shimomura, 2008 야마구치창벌레 Family Holognathidae Thomson, 1904 주걱벌레붙이과
Genus Cleantioides Kensley & Kaufman, 1978 주걱벌레붙이속 53. Cleantioides poorei Kwon & Kim, 1992 털보주걱벌레붙이 54. Cleantioides rotundata (Kussakin, 1982) 오목꼬리주걱벌레붙이 Family Idoteidae Samouelle, 1819 주걱벌레과
Genus Cleantiella Richardson, 1912 갯주걱벌레속
**55. Cleantiella n. sp.
56. Cleantiella isopus (Miers, 1881) 갯주걱벌레 Genus Idotea Fabricius, 1798 주걱벌레속 57. Idotea metallica Bosc, 1802 떠돌이주걱벌레 Genus Paridotea Stebbing, 1893 톱니주걱벌레속
１８ 58. Paridotea robusta Nunomura, 1985 톱니주걱벌레 Genus Pentidotea Richardson, 1905
59. Pentidotea rotundata Richardson, 1909 새우말주걱벌레 Genus Pentias Richardson, 1094 거친다리주걱벌레속
**60. Pentias n. sp.
61. Pentias thompsoni Collinge, 1916 넓적거친다리주걱벌레 Genus Synidotea Harger, 1878 둥근주걱벌레속
62. Synidotea hikigawaensis Nunomura, 1974 하키가와둥근주걱벌레 63. Synidotea laevidorsalis (Miers, 1883) 둥근주걱벌레
64. Synidotea nipponensis Nunomura, 1985 일본둥근주걱벌레 Suborder Asellota Latreille, 1802 물좀아목
Superfamily Janiroidea Sars, 1897
Family Janiridae Sars, 1897 물좀벌레과 Genus Ianiropsis Sars, 1897 수염갯가좀속
65. Ianiropsis koreaensis Jang & Kwon, 1990 둥근꼬리갯가좀 66. Ianiropsis serricaudis Gurjanova, 1936 긴턱갯가좀 67. Ianiropsis tridens Menzies, 1950 털보갯가좀 Genus Janiralata Menzies, 1951 두갈래바다좀속
*68. Janiralata sagamiensis Shimonura, 2006
69. Janiralata koreaensis Jang, 1991 매끈이마바다좀 Family Joeropsididae Nordenstam, 1933
Genus Joeropsis Koehler, 1885
**70. Joeropsis n. sp. 1
**71. Joeropsis n. sp. 2
Superfamily Stenetrioidea Hansen, 1905
１９ Family Stenetriidae Hansen, 1905
Genus Stenetrium Haswell, 1881
*72. Stenetrium sp.
(*, new to Korean fauna; **, new to science)
4.2. Descriptions of species
Subphylum Crustacea Pennant, 1777 갑각아문 Class Multicrustacea Latreille, 1802 연갑강
Superorder Peracarida Calman, 1904 낭하상목 Order Isopoda Latreille, 1817 등각목
Diagnosis. Body generally dorsoventrally flattened, occasionally cylindrical; carapace absent;
cephalon with 2 pairs of antennae, 1 pair of mandibles, 2 pairs of maxillae, and 1 pair of maxillipeds;
pereonites 7-segmented, usually free; pleonites 5-segmented, variously fused. Eyes sessile. Antennae uniramous, sometimes with rudimentary scale on antenna. Mandible normally with 1−3-articled palp;
incisor, molar, and lacinia mobilis usually present. Maxilliped composed of palp, endite, and epipod;
articles of palp variously fused depend on taxa. Pereopods uniramous, normally consisted of 7 pairs;
pereopod 7 developing with consecutive molts. Pleopods biramous, composed of 5 pairs; pleopod 2 of male with stylet on inner margin of endopod. Uropod biramous, consisted of 1 pair. Female with brood pouch or marsupium derived from coxal plate.
Key to suborders or infraorder of order Isopoda in Korean waters
1. Uropod operculiform; pleopods covered by uropod ··· Valvifera - Uropod not as above; pleopods not covered by uropod ··· 2 2. Pleopods 1 or 2 modified into operculiform in female; uropod located at distal region of pleoteslon;
··· Asellota - Pleopods 1−5 laminar or membranous in both sex; uropod positioned at lateral or ventral region ··
··· 3
２０
3. Mandibular molar process absent; maxillipedal endite expanding to distal region of palp article 4 ··· Limnoriidea - Mandibular molar process present (sometimes absent in several taxa); maxillipedal endite not reaching to distal region of palp article 4 ··· 4 4. Pleonites composed of 1 or 2 free pleonites; pleotelson vaulted; mandibular lacinia mobilis present ···Sphaeromatidea - Pleonites consisted of 5 free pleonites (some pleonites fused depend on species); pleotelson flat;
mandibular lacinia mobilis absent ··· Cymothoida
Suborder Cymothoida Wägele, 1989 부채벌레아목
Diagnosis. Body dorsoventrally depressed, sometimes cylindrical; pleonites 1−5 free (occasionally fused in some species); pleotelson, underside flat. Mandibular lacinia mobilis absent; molar process flat blade or conical. Maxillipedal endite barely longer than palp article 1 or absent. Pleopods 3−5 oval or oblong; distal end rounded.
Key to superfamilies of suborder Cymothoida in Korean waters
1. Body elongate and cylindrical; length 6 times longer than wide; uropodal exopod attached to proximal and dorsal region of protopod ...Anthuroidea - Body dorsoventrally depressed; length less than 6 times longer than wide; uropodal exopod attached to distal region of protopod... 2 2. Parasitic on crustaceans; mouthparts highly reduced ... Bopyroidea - Free-living or parasitic on fishes; mouthparts not as above ... Cymothoidea
Superfamily Anthuroidea Leach, 1914 마디벌레상과
Diagnosis. Body elongate and cylindrical. Antennule with 3 peduncular articles. Antenna with 5 peduncular articles. Mandibular molar process conical; palp consisted of 1−3 articles. Maxillule consisted of reduced inner lobe. Maxilla rudimentary. Maxilliped with diversely fused palp articles.
２１
Pereopods 1 or 1−3 subchelate. Pleonites 1−5 free or fused. Pleotelson with 0−2 statocysts dorsally.
Uropodal exopod positioned on proximal and dorsal region of protopod.
Key to families of superfamily Anthuroidea in Korean waters
1. Pleonites 1−5 much longer than wide; pleopod 1 not operculiform; pereopods 1−3 subchelate ··· Hyssuridae - Pleonites 1−5 as long as or shorter than wide; pleopod 1 operculiform; pereopod 1 subchelate ··· 2 2. Mouthparts not produced anteriorly; pleotelson with 1 pair of statocysts ··· Anthuridae - Mouthparts acutely produced anteriorly; pleotelson without statocysts ··· Paranthuridae
Family Anthuridae Leach, 1814 마디벌레과
Diagnosis. Body 10−15 times longer than wide. Pleonites less than twice as long as wide, variously separated by folds. Flagellum of antenna fewer than 10 articles, shorter than peduncles of antenna.
Mouthparts not produced anteriorly. Maxillipedal endite various in length: reaching palp article 3, obscure, absent; palp article variously fused. Pereopod 1 subchelate. Pleopod 1, only exopod operculiform. Pleotelson with 1 pair of statocysts.
Key to genera of family Anthuridae in Korean waters
1. Pleonites 1−3 separated by folds dorsally··· Amakusanthura - Pleonites 1−5 not separated by folds dorsally ··· 2 2. Flagellum of antenna consisted of 2 articles, mandibular palp 2-segmented ··· Caenanthura - Flagellum of antenna consisted of 3−4 articles, mandibular palp 3-segmented ··· 3 3. Maxillipedal palp consisted of 2 articles, carpus of pereopods 4−7 without stout seta ventrally ··· Cyathura - Maxillipedal palp consisted of 3 articles, carpus of pereopods 4−7 with stout seta ventrally ··· 4 4. Body without dark pigment; maxillipedal endite present or absent ··· Apanthura - Body with dark pigment pattern; maxillipedal endite absent ··· Mesanthura
２２ Genus Amakusanthura Nunomura, 1977
Type species. Amakusanthura longiantennata Nunomura, 1977.
Diagnosis. Body not darkly pigmented. Eyes present or absent. Pleon longer than wide; pleonites 1−3 separated by folds; pleonites 4−5 not separated by folds. Antennule of male with more than 10 flagellar articles. Mandibular palp 3-articled. Maxillipedal endite filamentous, triangular, or absent;
palp 3-articled; palp articles 1−2 fused, 3 free, 4−5 fused. Pereopod 1, propodal palm with step or prominent tooth. Pereopods 4−7, carpus triangular or trapeziform, with 1 stout seta on posterodistal angle. Uropodal endopod subequal to protopod in length; exopod with sinuous dorsal margin.
1. Amakusanthura n. sp. (Figs. 6−8)
Material examined. Holotype: Jeju-do: Non-ovigerous ♀, 5.5 mm, Beophwan-dong, Seogwipo-si, 33°13′36″N, 126°32′12″E, 26 April 2018, depth 33 m.
Paratypes: 4♀♀, same location as the holotype; 2♀♀, same locality as previous, 1 February 2018;
2♀♀, Seohong-dong, 33°13′48″N, 126°33′06″E, 26 April 2018, depth 51 m; 1 juvenile ♀, Donghong-dong, 33°13′48″N, 126°34′36″E, 26 April 2018, depth 59 m.
Description. Holotype, female: Body (Fig. 6A, B) 5.5 mm in length, 13 times longer than wide, smooth and slender. Cephalon square to globular, 0.8 times as long as pereonite 1; rostrum as long as anterolateral lobes; eyes very small; color not pigmented, white. Pereonites rectangular; pereonites 1−6 similar to each other in length; pereonite 7 about 0.7 times as long as pereonite 6; coxal plates of pereonites 4−6 visible dorsally. Pleonites visible dorsally; pleonites 1−3 separated by folds;
pleonites 4−5 fused dorsally, with partial lateral sutures; pleonite 6 free, with dorsal notch posteriorly.
Antennule (Fig. 6C) composed of 3 peduncular and 3 flagellar articles, sequentially slender in width. Peduncular article 1 rectangular, with 1 simple and 1 penicillate seta distally; article 2 subsquare, 0.6 times as long as article 1, with 3 simple and 4 penicillate setae distally; article 3 rectangular, 1.2 times longer than article 2, with 3 simple setae distally. Flagellar article 1 0.4 times as long as wide, with 1 simple and 1 penicillate seta; article 2 elongated, about 4 times longer than article 1, without seta; article 3 minute, square, with 7 simple setae distally.
２３
Fig. 6. Amakusanthura n. sp., holotype, female. A, Habitus, dorsal view; B, Habitus, lateral view; C, Antennule; D, Antenna; E, Mandible; F, Maxilla; G, Maxilliped. Scale bars: A, B = 1 mm, C−G = 0.1 mm.
２４
Anteanna (Fig. 6D) consisted of 5 peduncular and 4 flagellar articles, with geniculate peduncular articles between articles 2 and 3. Peduncular article 1 shorter than other articles, with 2 simple setae laterally; article 2 1.8 times longer than article 1, grooved to accommodate antennule, with 1 short simple seta laterally and 1 simple seta distally; article 3 0.7 times as long as article 2, with 3 simple setae distally; article 4 square, subequal to article 3, with 6 simple setae and 1 penicillate seta distally; article 5 elongate, 1.4 times longer than article 4, with 13 simple setae on both lateral margin and 1 penicillate seta on distal end. Flagellar articles much shorter than peduncular article 5, consecutively shortened; article 1 with 4 simple setae distally; article 2 with 1 simple seta; article 3 without setae; article 4 with 7 simple setae; articles 3 and 4 minutes.
Mandible (Fig. 6E), incisor with 1 prominent cusp and serrated margin; molar process bluntly rounded; palp consisted of 3 articles; article 2 1.5 times longer than article 1, slightly stouter than other articles, with 2 simple setae distally; article 3 shortest in length, with 4 serrate setae and a row of spines laterally, 1 serrate seta distally.
Maxilla (Fig. 6F) with 5 teeth distally and several fine setae laterally.
Maxilliped (Fig. 6G), endite extending to proximal end of first palp article, with 1 short simple seta apically; palp consisted of 3 articles; palp articles 1−2 fused, with 1 simple seta distally; article 3 free, with 2 simple setae laterally and 2 simple setae distally; articles 4−5 fused, oblique, smaller than other articles, with 4 simple setae laterally.
Pereopod 1 (Fig. 7A) prehensile, robust; basis continuously stouter distally, with 3 simple and 3 penicillate setae on anterior margin; ischium slender than basis, rectangular, with 1 simple seta on posterior distal end; merus much shorter than ischium, wider than long, with 1 simple seta on anterior and posterior distal end respectively; carpus triangular, with rough margin posterodistally and 8 simple setae along with posterior margin; propodus swollen, stepped on palm, with 12 simple setae on posterior margin and 3 simple setae on distal end; dactylus with 5 simple setae distally, a row of spines and 1 simple seta laterally; unguis as long as dactylus, much longer than those of other pereopods, with small accessory unguis. Pereopods 2−3 (Fig. 7B, C) similar to each other; basis elongate, oval, with 1 simple seta on posterior margin distally; ischium slightly shorter than basis,
２５
Fig. 7. Amakusanthura n. sp., holotype, female. A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7. Scale bar = 0.1 mm.
２６
with 2 simple setae on both lateral margins; merus about 0.5 times as long as ischium, tapering proximally, with several simple setae on posterior and anterior margins; carpus triangular, with several simple setae on posterodistal angle and fine setae along posterior margin; propodus elongate and oval, with simple setae on both lateral margins and 1 stout seta on posterodistal angle; dactylus with simple setae distally; unguis 0.5 times as long as dactylus, with small accessory unguis.
Pereopods 4−6 (Fig. 7D−F) subequal to pereopods 2 and 3 except for carpus; carpus more or less rectangular, with 1 stout seta on posterodital angle (absent in pereopod 5); propodus with several fine setae along both lateral margins and 1 stout seta on posterodistal angle. Pereopod 7 (Fig. 7G), subequal to previous 3 pereopods; carpus and propodus with dentate margin baring simple setae along posterior margin.
Pleopods (Fig. 8A−E) sequentially shortened in length; rami with plumose setae distally;
exopod slightly longer than endopod; pleopod 1 (Fig. 8A) operculiform, exopod about twice as long as wide; endopod 0.4 times wider than exopod; pleopods 2−5 (Fig. 8B−E), similar to each other, protopod with 1−2 simple setae on inner or outer margin.
Uropod (Fig. 8F, G), protopod rectangular, 2.4 times longer than wide, with plumose setae both lateral margins; exopod oval, almost 2 times longer than wide, surrounded by plumose and simple setae, with pointed apex; endopod sinuous distally, with plumose and simple setae along both lateral margins.
Pleotelson (Fig. 8H) 2.2 times longer than wide, concave proximally, convex medially, tapering distally, with 2 statocysts on dorsal surface proximally; apex with 5 simple setae; distal region with several simple setae on dorsal surface and lateral margins.
Habitat. Subtidal zone of sand flat.
Remarks. There have been reported 42 species of the genus Amakusanthura Nunomura, 1977 from all over the world (Boyko et al., 2008). Amakusanthura n. sp. is most similar to four species, Amakusanthura magnifica (Menzies & Frankenberg, 1966); Amakusanthura pimelia (Poore & Lew Ton, 1985); Amakusanthura paramagnifica Müller, 1992; Amakusanthura tengo Müller, 1992, by the following characteristics: (1) pleonites 1−3 are separated by folds, while pleonites 4−5 fused
２７
Fig. 8. Amakusanthura n. sp., holotype, female. A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5; F, Uropodal exopod; G, Uropodal endopod; H, Pleotelson. Scale bar: A−H
= 0.5 mm.
２８
dorsally; (2) the uropodal exopod has sinuous and acute distal region; (3) the pleotelson is concave laterally and tapering distally; (4) the maxillipedal endite is presence; (5) the propodus of pereopod 1 has a stepped palm. However, this Korean new species can be distinguished from them by having maxillipedal endite (lacking in the A. tengo), protruded inferior distal angle on carpus of pereopod 1 (not protruded in the A. pimelia), maxillipedal endite reaching to distal end of first palp article (over in the A. magnifica; not reaching in the A. paramagnifica) (Schultz, 1969; Poore & Lew Ton, 1985;
Kensley & Schotte, 1989; Müller, 1992).
２９ Genus Apanthura Stebbing, 1900 길쭉마디벌레속 Type species. Apanthura sandalensis Stebbing, 1900.
Diagnosis. Body not darkly pigmented. Pleonites as long as wide in length; pleonites 1−5 fused.
Antennule of males with less than 10 flagellar articles. Mandibular palp 3-articled. Maxillipedal endite triangular; palp 3-articled; palp articles 1−2 fused, 3 free, 4−5 fused. Pereopod 1, propodal palm with step or prominent tooth. Pereopods 4−7, carpus pentagonal, with 1 stout seta on posterodistal angle.
Key to known species of genus Apanthura in Korean waters
1. Eyes absent ··· Apanthura n. sp.
- Eyes present ··· A. koreaensis
2. Apanthura n. sp. (Figs. 9−11)
Material examined. Holotype: Jeju-do: Non-ovigerous ♀, 5.4 mm, Gapa-ri, Daejeong-eup, Seogwipo-si, 33°06′54″N, 126°16′42″E, 31 January 2018, depth 71 m.
Paratypes: 2♀♀, same location as the holotype; 1♀, Gapa-ri, 33°10′00″N, 126°14′48″E, 28 April 2018, depth 36 m.
Description. Holotype, female: Body (Fig. 9A, B) smooth and slender, 5.4 mm in length, 11 times longer than wide. Cephalon square, 0.7 times as long as pereonite 1; rostrum extending as long as anterolateral lobes; eyes absent. Pereonites oblong; pereonites 1−3 similar in length; pereonites 4−6 longer than previous three pereonites, subequal each other in length; pereonite 7 shorter than other pereonites, 0.7 times as long as pereonite 6; coxal plates of pereonites 5−7 visible dorsally. Pleonites 1−5 fused; pleonite 6 free, with middorsal notch posteriorly.
Antennule (Fig. 9C) consisted of 3 peduncular and 3 flagellar articles. Peduncular article 1 oblong, longer than wide, with 3 penicillate setae and 1 simple seta along lateral margin; article 2 square, about 0.5 times as long as article 1, with 3 penicillate setae on distal end; article 3 similar to article 2 in length, slender than article 2 in wide, with 1 penicillate seta and 3 simple setae distally.
